{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T19:16:48Z","timestamp":1790623008182,"version":"4.1.0"},"reference-count":60,"publisher":"Wiley","issue":"3","license":[{"start":{"date-parts":[[2020,5,15]],"date-time":"2020-05-15T00:00:00Z","timestamp":1589500800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Journal of Anatomy"],"published-print":{"date-parts":[[2020,9]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>The cranial ossification sequence in <jats:italic>Pleurodeles waltl<\/jats:italic> is widely used in phylogenetic analyses of amphibian origin and evolution. However, the patterns published to date are far from completely resolved and contain certain discrepancies. Based on a large sample of <jats:italic>P.\u00a0waltl<\/jats:italic> specimens ranging from early post\u2010hatching larvae to post\u2010metamorphic newts, we determined the most common cranial ossification sequence and revealed its intraspecific variations. Since thyroid hormones (THs) are involved in the mediation of skull development in salamanders, we studied the role of THs in the cranial development of <jats:italic>P.\u00a0waltl<\/jats:italic>. The normal sequence and timing of bone appearance were compared with those in larvae reared under conditions of high (in 1 and 2\u00a0ng mL<jats:sup>\u20101<\/jats:sup> triiodothyronine) and low [in 0.02% thiourea (TU), which inhibits thyroid gland activity] TH levels. Metamorphosis was greatly accelerated in the TH\u2010treated larvae and was arrested in the TU\u2010treated larvae, which retained the larval pattern of the palate and rudimentary external gills even after 2 years of the experiment. Early\u2010appearing bones (the coronoid, vomer, palatine, dentary, squamosal, premaxilla, parasphenoid, pterygoid, prearticular, vomer, frontal, parietal, exoccipital, in this order) arise at the same stages and ages, and follow the same ossification sequence under different TH levels. The timing of the appearance of bones normally arising in the late larval and metamorphic periods (the quadratojugal, orbitosphenoid, prootic, maxilla, nasal, os thyroideum, prefrontal, quadrate, in this order) changes depending on the TH level. The maxilla and nasal display the most pronounced reaction to changes in the TH level: they appear precociously in TH\u2010treated animals, while their appearance is postponed and they remain rudimentary in TU\u2010treated animals. Because of different responses to THs, the order in which late\u2010arising bones appear changes depending on the TH level. Although bones appearing early in larval ontogeny (e.g. the premaxilla, vomer, squamosal, palatine) display no TH\u2010induced reaction when they start to develop, their further differentiation shows dependence on THs, and these bones become TH\u2010inducible closer to metamorphosis. These findings indicate that TH involvement in the mediation of cranial development changes from minimal (if at all) in its early stages to maximal during metamorphosis. It is likely that the appearance of bones early in development is mediated by factors other than THs. Their further development is accompanied by changes in the mechanisms mediating their morphological differentiation. That is, likely non\u2010hormonal mediation becomes replaced or\/and complemented by hormonal mediation. The constituent parts of the same bone may exhibit differences in their reactions to changes in TH levels. Although in normal development, the overall cranial ossification sequence is constant, there was variation in the order in which late\u2010appearing bones was recorded. These observations suggest that this variation results from individual variability in the internal TH level. Comparison with other salamanders suggests that (a) the pattern of TH mediation described in <jats:italic>P.\u00a0waltl<\/jats:italic> is common for cranial development of metamorphosing urodeles and (b) the same bone may differ in its TH dependence in different salamanders, e.g. there are interspecific variations in the degree of TH dependence of individual cranial bones.<\/jats:p>","DOI":"10.1111\/joa.13210","type":"journal-article","created":{"date-parts":[[2020,5,16]],"date-time":"2020-05-16T03:09:56Z","timestamp":1589598596000},"page":"543-555","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":12,"title":["Skull development in the Iberian newt, <i>Pleurodeles waltl<\/i> (Salamandridae: Caudata: Amphibia): timing, sequence, variations, and thyroid hormone mediation of bone appearance"],"prefix":"10.1111","volume":"237","author":[{"given":"Sergei V.","family":"Smirnov","sequence":"first","affiliation":[{"name":"A. N. 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